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(Piperaceae) from Uruguay Phytotaxa 244 (2): 125–144 ISSN 1179-3155 (print edition) http://www.mapress.com/j/pt/ PHYTOTAXA Copyright © 2016 Magnolia Press Article ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.244.2.2 Taxonomic revision of Peperomia (Piperaceae) from Uruguay PATRICIA MAI1, ANDRÉS ROSSADO2, JOSÉ M. BONIFACINO2,3 & JORGE L. WAECHTER4 1 Licenciatura en Gestión Ambiental, Centro Universitario de la Región Este, Universidad de la República, Rocha, Uruguay. 2 Laboratorio de Sistemática de Plantas Vasculares, Facultad de Ciencias, Universidad de la República, Montevideo, Uruguay. 3 Laboratorio de Botánica, Departamento de Biología Vegetal, Facultad de Agronomía, Montevideo, Uruguay. 4 Departamento de Botânica, Universidade Federal do Rio Grande do Sul, Porto Alegre, RS, Brasil. Corresponding author: [email protected] Abstract The genus Peperomia is represented by eight species in Uruguay: P. catharinae, P. comarapana, P. hispidula, P. increscens, P. pereskiifolia, P. psilostachya, P. tetraphylla and P. trineuroides. Peperomia psilostachya is reported for the first time for the flora of Uruguay, from material collected in moist hillside and riverside forests from the northeast and east of the coun- try. Three new synonyms are proposed: P. arechavaletae var. arechavaletae as synonym of P. trineuroides, P. arechavaletae var. minor of P. tetraphylla and P. trapezoidalis of P. psilostachya. Lectotypes for P. arechavaletae, P. arechavaletae var. minor and P. tacuariana, and a neotype for P. herteri are designated. The taxonomic treatment includes synonymies used in Uruguay, morphological descriptions, distribution and habitat data, phenology, conservation assesment, observations, and material examined for each species treated. A species identification key, plant illustrations and distribution maps in Uruguay are provided. Resumen El género Peperomia está representado en Uruguay por ocho especies: P. catharinae, P. comarapana, P. hispidula, P. incre- scens, P. pereskiifolia, P. psilostachya, P. tetraphylla y P. trineuroides. Peperomia psilostachya es registrada por primera vez para la flora de Uruguay a partir de material colectado en bosques de quebrada y ribereños del noreste y este del país. Se pro- ponen tres nuevas sinonimias: P. arechavaletae var. arechavaletae como sinónimo de P. trineuroides, P. arechavaletae var. minor de P. tetraphylla, y P. trapezoidalis de P. psilostachya. Se designan lectotipos para P. arechavaletae, P. arechavaletae var. minor y P. tacuariana, y un neotipo para P. herteri. El tratamiento taxonómico incluye para cada especie sinonimias usadas en Uruguay, descripciones morfológicas, datos de distribución y hábitat, fenología, evaluación del estatus de conser- vación, observaciones y material examinado. Asimismo, se presenta una clave para la identificación de las especies, ilustra- ciones y mapas de distribución en Uruguay. Key words: identification key, geographic distribution, conservation assessment, new records, new synonyms, typification Introduction Peperomia Ruiz & Pav. (Piperaceae) is one of the largest genera of angiosperms (Frodin 2004), including 1432 species (Mathieu et al. 2015) of terrestrial, epipetric or epiphytic herbaceous plants (Guimarães et al. 1984). The genus has a pantropical distribution, with the highest diversity in the Neotropics (Wanke et al. 2006, Mathieu et al. 2015). In America the genus occurs from Bermuda and the southeastern United States (Boufford 1982) to southern Chile, central and littoral Argentina and southern Uruguay (IBODA 2015). Several morphological synapomorphies support this genus, such as a single carpel, unitegmic ovule, 16-nucleate embryo sac, two monothecal bisporangiated stamens, and small non-aperturate pollen (Tucker et al. 1993, Jaramillo et al. 2004, Wanke et al. 2006). The monophyly of Peperomia is supported by molecular and morphological studies (Jaramillo et al. 2004, Wanke et al. 2006) but the traditionally used infrageneric classification, as proposed by Dahlstedt (1900), has shown some incongruency (Samain et al. 2007). Recently a new infrageneric classification was proposed based on morphological and molecular data (Frenzke et al. 2015), revealing fourteen monophyletic groups and therefore solving the previous problems regarding the infrageneric classifications. Accepted by Zhi-Qiang Zhang: 1 Jan. 2016; published: 18 Jan. 2016 125 Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0 The taxonomy of Peperomia is rather complex and controversial (Mathieu & Callejas 2006, Samain et al. 2011, Zanotti et al. 2012), mainly because the genus has been poorly collected and studies have been largely based on dry herbarium material with distinctive architectural features either missing or deformed (Mathieu & Callejas 2006). These circumstances have led to a confusion in the taxonomy of closely related taxa and to an over-description of species, with numerous synonyms for a given taxon throughout its geographic range (Mathieu & Callejas 2006, Frenzke et al. 2015). Taxonomic revisions of Peperomia for Uruguay were published by Trelease (19411), Trelease & Herter (19522) and Marchesi (1968) who cited six taxa: P. arechavaletae A. C. P. de Candolle (1917: 466), P. arechavaletae var. minor Herter (1952: 45), P. blanda (Jacq.) Kunth var. pseudodindygulensis (C. DC.) Yuncker (1953: 218), P. catharinae Miquel (1843: 127), P. hispidula (Sw.) A. Dietrich (1831: 165) and P. reflexa (L. f.) A. Dietrich (1831: 180). Later on, P. comarapana A. C. P. de Candolle (1915: 8) was cited for western Uruguay by Alonso Paz (1989) and P. pereskiifolia (Jacq.) Kunth ([1815] 1816: 68) was cited for eastern Uruguay by Zuloaga et al. (2008), although the former study does not mention supporting voucher or reference bibliography. This work aims to update the taxonomic knowledge of Peperomia for Uruguay. It provides an identification key for Peperomia species occurring in the country and proposes new synonyms, three lectotypes and a neotype. For each taxonomic entity treated synonymies, morphological descriptions, geographic distribution, habitat, phenology information, maps and illustrations are included. Material and methods The taxonomic revision was conducted based on herbarium specimens hosted at MVFA, MVFQ, MVHC, MVJB and MVM herbaria and specimens collected in the field. Additional specimens were examined from CTES (Argentina) as well as HAS and ICN (Brazil) (Thiers continuously update). A set of herbarium specimens from different geographical zones of Uruguay was selected and analyzed for each species. Fieldwork was carried out from September 2012 to November 2013 throughout the country. Collected material was deposited at MVJB and was identified according to the following literature: Yuncker (1953), (1958), (1974), Marchesi (1968), Novara (1998), Monteiro & Guimarães (2008) and Zanotti et al. (2012). A key for the identification of species was built based on morphological characters observed in herbarium material and in plants growing in situ or ex situ. Additionally, an electronic key using Lucid 3.5 (www.lucidcentral.org) was generated and can be accessed at http://www.thecompositaehut.com/www_tch/webinvestigacion/clave_virtual_flora/ cvfu_claves.html. The taxonomic treatment includes synonyms used within Uruguay, morphological descriptions, geographic distribution with emphasis in the Southern Cone of South America, habitat, phenology, assessment of conservation status, morphological and taxonomic observations, specimens examined, illustrations and distribution maps within Uruguay. General geographic distribution was obtained from several sources (Monteiro & Guimarães 2008, Zanotti et al. 2012, GBIF 2015, Tropicos 2015) and complemented by the herbarium specimens analyzed. We added detailed information for the Southern Cone, which includes Argentina, southern Brazil (Paraná, Santa Catarina and Rio Grande do Sul), Chile, Paraguay and Uruguay. Geographic distribution information of Brazil was taken from Guimarães et al. (2015) and for Argentina, Chile and Paraguay from herbarium specimen labels and the “Flora del Cono Sur” database (IBODA 2015). Geographic distribution within Uruguay was obtained from data present in herbarium material and from specimens collected during fieldwork. Habitat categories were adapted from Alonso Paz & Bassagoda (2002) and Brussa & Grela (2007) as follows: hydrophilous forest (‘bosque hidrófilo’) occurs in temporally flooded areas dominated by hydrophytes and generally the characteristic species is Erythrina cristagalli L.; open thorn forest (‘bosque parque’), a type of wooded savanna dominated by Aspidosperma quebracho-blanco Schltdl., Prosopis affinis Spreng., Prosopis nigra (Griseb.) Hieron. and Vachellia caven (Molina) Seigler & Ebinger, among other species; psammophilous forest (‘bosque psamófilo’) occurs in sandy soils in coastal areas; riverside forest (‘bosque ribereño’) occurs in floodplains of rivers and streams; xeric hillside forest (‘bosque serrano’) occurs on hill slopes (includes rocky escarpment and shrublands) and moist 1 Trelease’s work was firstly published in 1939, but due to the Second World War was disseminated in 1941. The taxa published in this work have 1941 as publication year, according to Art. 31.1 of ICB (2012). 2 Trelease & Herter appear as the authors of “Piperaceae Uruguayenses Curae Posteriores” in 1952. However Trelease died in 1945, therefore we consider Herter as the author of the new entities proposed in this publication. 126 • Phytotaxa 224 (2) ©
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